Arachnologische Mitteilungen43:66-78 Nuremberg,July2012 Assemblages of herb-dwelling spiders (Araneae) ofvarious steppe types in Ukraine and the Central Chernozem region of Russia NinaY.Polchaninova doi;10.5431/aramit4312 Abstract; A total of 107 spider species from 15 families were recorded in the herbaceous vegetation ofthe steppeecosystemsofUkraineandtheCentralChernozemregionofRussia.Araneidae,Thomisidae,Salticidaeand Theridiidaewerethe mostspecies-rich.The speciescomposition depended on the steppetype;adjacentforest habitatsinfluencedsteppefaunaintheforest-steppeandnorthernpartofthesteppenaturalzone.Thenumberof generalist,forestandwetlanddwellingspeciesinthesteppevegetationshowedatendencytodecreasetowards thesouth.Dominanceofherb-dwelling spiderswasspecifictoeach steppetype;nosinglespecieswasfoundto predominateinallthesteppe habitats. Keywords:dominancestructure,speciesdistribution,spidercommunities,steppeecosystems Steppes are the most transformed ecosystem in assemblages(CHERNOV1975).Habitatpreferenceof Ukraine.The steppe natural zone comprises 40% of speciesdependsonthenaturalzone(KÜHNELT1943, the countryandabout 80% ofthis territorywas once Walteri960,Bei-BienkO 1966). Accordingtothe covered with steppe vegetation. Presently, only 3% so-called principleofzonalchangeofhabitats’(BEI- ofrelativelyundisturbedvirginsteppeshavesurvived BIENKO 1966),ortheprincipleof‘relativestenotopy’ intact.They are preserved mainly in nature reserves (Schaefer1992),widespreadspeciesmovingnorth- or on gully slopes and saline lands not suitable for wardscanchangetheirhabitatstodryerwarmeropen agriculture (KOTENKO 1996). siteswithsparsevegetation,whilegoingsouthwards, Allsteppereservesareisolatedtovariousdegrees, thesamespeciesinhabitmoisterandshadierhabitats suchthattheycanbeconsideredasecologicalislands, with dense vegetation cover.These two phenomena surrounded by agricultural landscapes (MALYSHEV wereillustratedbytheexampleofspidercommunities 1980). Different components of the steppe biota ofthe Urals transect (ESYUNIN 2009), oakforests of respond to isolation in different ways (KOTENKO the East European Plain (ESYUNIN et al. 1994), and 1996,Hanser&cHuntly2006,Laiolo&Tella Ukrainian steppes (POLCHANINOVA 1990a, 1996). 2006).Spidersmovequiteeasilyfromnaturalhabitats Investigation ofthe spider fauna and ecology in the to agricultural fields, however many ofthem do not area in question began in the 1980s in the Central penetratebeyond the field margins. Crops are popu- ChernozemReserveofRussia(PiCHKA1984 ,1984b) latedmainlybyeurytopicspeciestypicalofmeadows and in the steppe reserves ofUkraine (POLCHANI- ordisturbedhabitats(SEYFULINA&TCHERNYSHEV NOVA 1988, 1990b). Based on the data obtained, a 2001, SEYFULINA2010). firstattemptwasmadetoanalyzethefaunaandspider Adjacent habitats have an undeniable impact on communities of the steppe ecosystems of Ukraine steppe communities (CHERNOV &c PENEV 1993), (Polchaninova 1990b). Further research focused especially in the forest-steppe zone where forest on all steppe reserves in Ukraine (GURIANOVA &c and steppe are both zonal plant formations, and Khomenko 1991, Polchaninova 1998, Pol- an active species interchange can be supposed to chaninova &C PROKOPENKPO 2007a, PROKO- take place. To the north and to the south the dif- PENKO 2001, Prokopenko et al. 2008) and the ference in microclimatic conditions between zonal adjacent territoryofRussia (POLCHANINOVA2003, and intrazonal habitats increases, which results in 2009, Ponomarev 2005, 2010). Thus, it became higher species specificity oftheir animal and plant possibletoanalysethespeciescompositionanddomi- nance structure ofspiderassemblages to seewhether NinaY.POLCHANINOVA,DepartnnentofZoologyandAnimalEcology, theydependonthe steppe type.Thepresentpaperis KharkivNationalUniversity,4,SvobodySq.61022Kharkiv,Ukraine. partofacomprehensivestudyofspidercommunities E-Mail:[email protected] ofsteppeecosystemsofUkraineandEuropeanRussia submitted:/.12.2011,accepted:4.6.2012;onlineearly:31.7.2012 and concerns herb-dwelling spiders only. Spidersinsteppein UkraineandCentralChernozemRussia 67 Materialsandmethods The material was collected from three localities in the Central Chernozem region ofRussia, and eight localities in the Left-bankUkraine (Fig. 1). Central ChernozemisasegmentoftheEurasian chernozem (blacksoilcontainingahigh percentageofhumus)beltthatlieswithin the EastEuropean Plain (also known as the RussianPlain).Itisawell-delimited geographical and administrative region. The Left-bank Ukraine in the present contextis a territorythat stretches from the leftbankofthe Dnieper River east- ward to the border ofthe state. We investigated five types of frag- ments of virgin steppes in the East European Plain (classification accord- ing to Gribova et al. 1980). Meadow, forb-bunchgrassandbunchgrasssteppes belong to the categoryofzonal steppes, whilesandyandchalkysteppesbelongto theazonalsteppes.InRussiangeobotany, meadow steppes situated in the forest steppe natural zone are called ‘northern Fig.1:Mapoflocalities(I-forest-steppezone,II-steppezone:lla-subzone steppes’ in contrast to genuine steppes, offorb-bunchgrasssteppes,Mb-subzoneofbunchgrasssteppes.For which form two subzones ofthe steppe numberoflocality,seeTab.1). natural zone (Fig. 1,Tab. 1).We regard sandy steppes situated in different steppe subzones Tab. 1.Hereandadditionallyinthegraphsandtables, separately as northern and southern variants. The thelocalitiesarearrangedfromnorthtosouth.Names studied localities, investigation period and abbrevia- ofsettlementsandnaturereservesaretranslatedfrom tionsusedfurtherinthetablesandgraphsaregivenin Russian and Ukrainian respectively. Tab.1:Listandcharacteristicsofstudysitesincludingtypeofsteppehabitat. Naturezone/ Nameoflocality Steppetype Yearsof sitearea abbrev. Region subzone study (ha) 1. Streltsovskaya meadowsteppe 1998 730 meadl KurskArea,Central steppe 1999 Chernozem Res. 2007 5F36'N 36U2T m 248 a.s.l. 2. Kazatskaya meadowsteppe 1999 1010 mead2 KurskArea,Central steppe 2000 Chernozem Res. 2007 5T32'N 36°20T m Forest-steppe 230 a.s.l. zone 3.Yamskaya meadowsteppe 2001 500 mead3 BelgorodArea,Belogorye steppe 2002 Res. 51TLN3r45T m 193 a.s.l. 4.Mikhailivska meadowsteppe 1985 160 mead4 SymskaAreaUkrainian Tsilyna 1986 steppe Res. 1988 50°34'N 44°12T m 172 a.s.l. 68 N.Polchaninova Naturezone/ Nameoflocality Steppetype Yearsof sitearea abbrev. Region subzone study (ha) 5. Striltsivskyi forb-bunchgrass 1986 1000 forbl LuhanskArea,Luhanskyi step steppe 1988 NaturalRes. 2009 49°17'N 40°00T m 147 a.s.l. Steppezone: 6. Khomutivskyi forb-bunchgrass 1982 1000 forb2 DonetskArea,Ukrainian Northern step steppe 1983 steppe Res. subzone 2004 4ri7'N38°10'E m offorb- 57 a.s.l. bunchgrass 7. Striltsivskyi chalkysteppe 2009 80 chalk LuhanskArea steppe zakaznyk 2011 49°18'N 39°50T m 100 a.s.l. 8.Dnieprovsko- Northernsandy 1999 300 sandN DnipropetrovskArea Orilskyi Res. steppe 2000 48°30'N 3T45T m 2003 53 a.s.l. 9.Askania-Nova bunchgrass 1984 11000 bunch KhersonArea, Res. steppe 1985 46°28^N 33°58T m 1988 29 a.s.l. Steppezone: 10.Ivano- Southern sandy 1989 2074 sandSl KhersonArea, Southern Rybalchanskyi steppe 1990 Chernomorskyi Res. subzoneof 1991 46°27'N 32°07T m bunchgrass 6 a.s.l. steppe 11. Solenoozernyi Southernsandy 1995 1325 sandS2 KhersonArea, steppe 1996 Chernomorskyi Res. 1998 46°27'N3F58T m 1 a.s.l. Ineachlocality,investigationswerecarriedoutfor immature specimens could not be distinguished, we two consecutiveyears.The materialwas collectedby considered thegenus as awhole ingeneralquantita- sweeping with an entomological net (d=30 cm), as tive analysis, and then species relationships within wellasbyhandcollecting.Quantitativesampleswere thegenuswere estimatedseparatelybasedonmature takeneverymonthfromMaytoSeptember,fivesam- individuals. ples of50 sweeps each.The material obtained from Inordertodeterminerelativeabundance,weused two vegetation seasons was considered as a general theTischlerrating scale,where eudominant n>10%, sample. For comparative analysis, we chose plots in dominant 5<n<10%, subdominant 2.5<n<5%, rece- naturereserveswithstrictlyprotectedsteppe,because dent l<n<2.5%, and subrecedent n<l% (TISCHLER traditionalregime management such as haymowing 1949). We considered eudominants and dominants dramatically impacts upon species composition and together as a dominant complex. The ecological structure of spider communities (POLCHANINOVA groups ofspecieswere determinedbased on the data 2004). In order to obtain more information on the on their habitat preference within the forest-steppe spider species composition oflocal faunas and vari- andsteppezonesoftheEastEuropeanPlain(Tab.2). oussteppetypes,weconductedadditionalresearchin Wedefinegrasslandspeciesasspeciesoccurringinall differentyears, and also used the data from available habitatswithpredominanceofherbaceousvegetation, publications (PiCHKA 1984a, 1984b, GURYANOVA in our case meadows, steppes, and open slopes of &KHOMENKO 1991,Prokopenko2001).Intotal, gullies. We consider permanent residents ofone or morethan 12,000spiderspecimenswerecollected.A more steppe types to be typical species. Sometimes list ofspecies is given in the Appendix. Species are theymayoccurin other habitats as rare finds. arrangedbyfamilies accordingto PLATNICK (2012). Similarity of spider assemblages of the studied Withthemethodofnetsweeping,alargenumber habitatswasdeterminedbyclusteranalysisperformed ofjuvenilespiderswerecollected.Inmanycases,they inStatistica7(STATSOFTINC.2004).Twoyearquan- could be identified to species level as theybelonged titativesamples (seeabove)werepooledtogetherand to a single species ofthe genus in agiven locality, or apercentageofeachspeciesofthetotalsampleateach differed in period ofmaturity. For the generawhose sitewascalculated.WeusedtheWardsalgorithmasa Spidersinsteppein UkraineandCentralChernozemRussia 69 dustermethodandEuclidiandistanceas asimilarity steppe.The spider fauna ofmeadow steppe situated measure. in forest-steppe zone differs significantly from that ofsteppe habitats in steppe zone.Within the steppe Results zone,spidersaremoreevenlyspread,andthefaunaof Speciescompositionandspeciesrichness eachsteppetypehasonly7-12%ofspeciesnotfound A total of107 spider species from 15 families were in other steppe types. recorded in herbaceous vegetation at 11 study sites. Species richness of herb-dwelling spiders was Four families were the most species-rich: Araneidae maximal in both fragments of the forb steppe and (20species, 18.7%oftotalspeciesinstudysites),Sal- minimal in both fragments of the southern sandy ticidae (19 species, 17.8%),Thomisidae (18 species, steppe (Appendix). In the largest steppe fragment 16.8%), and Theridiidae (13 species, 12.1%). Only (bunchgrass steppe in Askania-Nova) the species Araneidaeweredistributedquiteevenly,rangingfrom richness was slightlylower than that in the smallest 22%ofthespeciesinforbsteppesto27%inthesandy one (chalkysteppe inMilove District). ones. Linyphiidae and Clubionidae occurred mainly inmeadowsteppes (Appendix).Thomisidae reached Ecologicalgroupsofspecies their maximum of species richness in bunchgrass Meadowsteppes arecharacterizedbyalarge number steppes (23%) and fell to a minimum in meadow offorestandwetland species (Tab.2). Some ofthem steppes (13%). Salticidae are known to increase in appear in the steppe in wet years, and nine species terms of the number of species towards the south are permanent residents. The number ofwetland (Nenilin 1984, Mikhailov 1997). In our collec- and forest species gradually decrease towards the tion,wefoundunexpectedlyfewsalticidspeciesinthe south. In different steppe types, generalist species bunchgrasssteppe(4species,10%);intheothersteppe comprised 15-20%.The numberofsteppe species at types this increase could be seen not in absolute but all sites was significantly lower than that ofspecies onlyin relative numbers because ofimpoverishment widelydistributedingrasslands. In meadowsteppes, ofthe speciescompositioninthe southernsteppes (9 we found no steppe species. species, 15% in meadow steppes, 11 species, 17% in Therewere no specialist species inthe herb layer forbsteppes,9species,20%innorthensandysteppes, ofsteppe vegetation. Eight species whichwere con- 7 species,21% in the southern ones). sideredtobetypicalwerealsofoundinotherhabitats; A third ofthe species found in meadow steppe however, they reached their maximal abundance have not been recorded southwards in genuine onlyin a certain steppe type.These species are Run- Tab.2:Ecologicalgroupsofspidersofvarioussteppetypes. Numberofspeciesinsteppetype Ecologicalgroups meadow forb-bunch- chalky sandyN bunch-grass sandyS steppe grass generalist 10 11 9 8 6 5 wetland + forest 17 7 2 4 3 1 forest 7 3 1 wetland 7 2 2 grassland + forest 5 10 9 9 5 6 edges grassland 9 12 10 11 9 8 steppe 8 6 2 6 2 unspecified 6 9 2 4 7 5 1 1 typical 2 1 1 1 3 70 N.Polchaninova 100% Others ^ 75% 03 Miturgidae q; Oxyopidae 1 0 Salticidae Theridiidae c 50% 1^ Philodromidae _o H Thomisidae S o Dictynidae QO. ^ Araneidae Q. 25% 0% sites Fig.2:Familyabundance(proportionoftotal numbersoffamiliesateachsite) inspiderassemblagesofherba- ceousvegetationofsteppehabitats.Sitesarearrangedfrom northtosouth. Among five steppe species of the southern steppesubzone,threespeciesoccurredinsandy and chalky steppes of the northern subzone whiletwo species did notextendbeyondzonal We boundaries. did notfindtendencies in dis- tribution ofthe other species; 22 species were found as singletons,thereforewe cannotjudge their habitat preference. Familyabundance The herbaceous vegetation ofsteppe habitats was dominated by Araneidae at all sites (Fig. 2). The second largest family Thomisidae preferredforb and chalkysteppes.The relative LinkageDistance numberofDictynidaewentdownfrommeadow Fig.3:Dendrogramofbiocoenoticsimilarityofherbdwellingspider steppestobunchgrassandsouthernsandyones. assemblagesofvarioussitesbasedonthe%ofeachspeciesin Philodromidae, likewise, were least abundant thetwoyearsamplesineachsiteusingWard'smethodascluster algorithmand Euclidiandistanceassimilaritymeasure. in the three southern sites while they reached a maximum in forb steppes and in the‘mead4’ ciniagrammica, Pellenus seriatus and Yllenus vittatus site.ThedistributionofSalticidaedidnotshowalati- in southern sandy steppe, Heliophanuslineiventris in tudinal trend.An unusuallylarge individual number southern sandyand bunchgrass steppe, and Uloborus ofthis familywas found in one ofthe sandy steppe A walckenariusin sandyand chalky steppes. sites. characteristicfeatureofthespiderassemblage Thirteengeneralistandgrasslanddwellingspecies ofthe bunchgrass steppewas the high abundance of occurred in almost all investigated steppe fragments Oxyopidae. (Appendix). Four species found in steppes of the forest-steppe and northern steppe subzone changed Biocenoticsimilarities A their habitat preference to forests orwetlands in the dendrogram ofbiocenotic similarity ofspider as- southern subzone.Tenwetland species spread to the semblagesofvarioussitesbringsthemtogetherbased souththroughsteppefragments nofurtherthan sites on a zonal-subzonal principle (Fig. 3). First, two ‘forbl’and‘sandyN’;southwardstheyalsomovedinto clusters ofthe sites ofthe forest-steppe and steppe intrazonal habitats. Five steppe species were widely zone are separated, and then the sites of northern spread in the steppe zone, but absent in meadow and southern subzones are divided into two groups. steppes; three species occurred in meadow steppes As expected, assemblages oftwo neighbouring frag- onlyinthesite‘mead3’.Itisthedriesteasternsitewith ments of southern steppes were similar, however, a character ofvegetation closer to genuine steppes. theirsimilaritywaslessmanifestthanthatofthetwo Spidersinsteppein UkraineandCentralChernozemRussia 71 adjacent meadow steppe plots and even ofthe forb grasslands,butinthesouthitisparticularlyabundant. and chalkyones. Phylloneta impressa, another dry grassland species, preferred sandy and chalky steppes. Dictyna arundi- Dominancestructure nacea,bycontrast,waseudominantinmeadowsteppes In the study areas, 13 species occurred in all steppe and occurred in high numbers in the forb ones.This habitats.Inaddition,twospecieswerefoundsporadi- speciesischaracteristicofgrasslandsandforestedges callyinbothnorthernandgenuinesteppes.However in the forest-steppe and northern part ofthe steppe none ofthem dominated in all steppe types. zone.Thegroupofspecieswithanarrowerdominance Neosconaadiantawas distinguishedbythewidest spectrumwasrepresentedhyNeottiurabimaculataand habitat spectrum (Tab. 3). It is common in various Araneusquadratusinmeadowsteppes,andbyThomisus Tab.3:Dominantspiderspeciesindifferentsteppehabitats. eudominantn>10%,^^-dominant,5<n<10%,W-subdominant2,5<n<5%, recedent<n<2,5%,#-subrecedentn<1%ofspiderscollected ineach locality Localities .apecies 1 2 3 4 5 6 7 8 9 10 11 • • Neottiurabimaculata • a • • • Araneusquadratus • • • Cheiracanthiumpunctorium a a • • • • • • • • • Evarchaspp. • • • • • • • • Dictynaarundinacea a a • • • • • • • • Tibellusspp. • • • Xysticusspp. • • • • • Agalenatearedi • • • • • • • • • • Mangoraacalypha • • • • • • • • • • • • • • • Neosconaadianta • • • • • • • • • • • • • Heliophanusspp. • • • • Phyllonetaimpressa • • • • • • • • • Cheiracanthiumpennyi • • • • • • • • • • Philaeuschrysops • • • • • • Thomisusonustus • • • Uloboruswalckenaerius • a a • • Oxyopesheterophthalmus • • • 72 N.Polchaninova Despite the local differences in species compositionanddominancestructureofspider assemblages,anumberofcharacteristicfeatures canbe distinguished foreverysteppe type.All the spider complexes ofsteppe herbage had a polydominant structure. In the four sites of meadow steppes, the group of dominants consisted of Dictyna arundinacea (16.0-24.3%), Tibellus oblongus (8.7-14.2%) and Xysticus cristatus (6.0-9.3%). Araneusquadratuswdcs,amemberofthiscomplex in three sites,NeottiurabimaculatUyLarinioides patagiatuSy Cheiracanthiumpunctorium,Evarcha arcuatay andNeosconaadiantawerelocaldomi- nants in notmore thanone site each.Marpissa pomatiay Hypsosinga sanguinea, and Singa hamata were subdominants in the meadow steppes. Southwards in genuine steppes they did not occur, or were found only as a few specimens.Howeverwecannotconsiderthem Fig.4:Relativeabundance(%ofindividualsoutofthetotal number tobetypicalspeciesofmeadowsteppesbecause ofspiderscollectedateachsite)of(A) Tibellusoblongusand T. theywere also numerous inwetlands (personal macellusand (B)XysticuscristatusandX.striatipesin herbaceous vegetationofdifferentsteppehabitats.Sitesarearrangedfrom observation). The specificity ofthe spider as- northtosouth. semblage ofthe driest eastern site ‘mead4’ is worthmentioning.TheabundanceofNeoscona onustus and Philaeus chrysops in genuine steppes. Six adiantaandtheappearanceofEvarchamichailoviand species were dominants at only one site {Uloborus Xysticusstriatipes make it similar to the assemblages walckenaerius, Cheiracanthiumpennyi^ C.punctorium, offorb steppes. Philodromus histrio^ Pellenesseriatus, and Yllenus vit- Xysticus striatipes (9.3-15.7%), Tibellus spp. tatus). (10.7-12.6%), andNeosconaadianta (8.0-9.7%) were Offurther interest are the latitudinal changes of common dominants offorb steppes,withAgalenatea abundance of common species of the same genus. redii and Mangora acalypha co-occurringwith them In the pairs Tibellus oblongus-macellus and Xysticus in the northern site, and Philaeus chrysopSy Evarcha cristatus-striatipesy the first species ofeach pair was michailoviyHeliophanusflavipesand Thomisusonustus A abundantintheforest-steppeandinthenorthofthe in the southernone. range ofspecies typicalofdry steppe zone (Fig. 3).The second species ofeachpair grasslands appear in these steppes {Heterotheridion appeared in genuine steppes, and in case ofXysticuSy nigrovariegatumy Theridion innoccuumy Heriaeus ob- replaced the first one. Further to the south they all longuSyTibellusmacelluSyXysticusmarmoratus,Synageles decreased in number or disappeared in steppe plots, hilarulusy and Philaeus chrysops). However, we found although they remained quite common in adjacent no species specific for the forb steppes only. forest stands (POLCHANINOVA 1990a). Heliophanus Thedominancecomplexofchalkysteppeincluded flavipesoccurredinaUsteppetypes.Inthethreesouth- Xysticus striatipes {2S.S%)yAgalenatea redii (11.6%), ern sites, H. lineiventrisco-occurredwith H.flavipes Neoscona adianta (10.0%), Tibellus spp. (6.4%), PhT andtogethertheymadeupasignificantproportionof laeuschrysops(6.7%),andDictynaarundinacea(5.8%). the individuals found in these sites (Tab. 3).Evarcha Inthe northern sandysteppeitconsistedofNeoscona arcuatawas common in meadow steppes and in the adianta(14.9%),Xysticusstriatipes(11.0%),Mangora northern sandy steppe. E. michailovi occurred in acalypha (8.9%), Dictyna arundinacea (5.8%), Phi- forb steppes andpenetratedinto one site ofmeadow lodromus histrio (7.7%), and Uloborus walckenaerius steppes. Southward in the subzone of bunchgrass (5.3%).Intheareainvestigated,thelattertwospecies steppes, neither ofthese specieswas found. areassociatedwithvegetationofsandysoil.Theywere also common in calcareous lands (POLCHANINOVA SpidersinsteppeinUkraineandCentralChernozemRussia 73 2010). Simitidionsimileand Cyclosaoculatapreferred bothhabitatandvegetationlayermovingfromherbs the same habitats but they had larger ecological to the ground (POLCHANINOVA 1990a).Agroup of flexibility, inhabiting meadows, shrubbysteppes and southern species which spread to the north through & even open deciduous forests (POLCHANINOVA the dry open sites made up only 4% ofthe species Prokopenko 2007b). Gibbaranea bituberculata and found in the steppe habitats. Our data confirm a G. ullrichicametothesteppeplotsfromneighbouring decrease ofgeneralist, forest and wetland dwelling pine forests. species to the south and predominance ofgrassland The spider assemblage ofthe bunchgrass steppe species in genuine steppes. The same tendencies wascharacterizedbylargenumbersofthreeeudomi- in distribution ofecological groups ofspiders were nants Neoscona adianta (29.8%), Oxyopes heteroph- shownbyESYUNIN (2009) in a transectthrough the thalmus (26.2%), and Thomisus onustus (14.0%), the Urals. occurrenceofonesubdominantXysticusstriatipes,and The relative abundance of spider families in single finds ofotherspecies. Southern sandysteppes steppe vegetation changes depending on the steppe were dominated atboth sitesbyHeliophanusflavipes^ type.Thisisconfirmedbydataonsteppelocalitiesin H. lineiventris(15.6-20.0%),Neosconaadianta(11.0- other regions.Thus, in mountain steppes ofBashki- 12.2%),andPhyllonetaimpressa(5.6-6.3%).Inoneof ria (Southern Urals), Tetragnathidae, Thomisidae, them(Ivano-Rybalchanskysite)thedominancecom- and Araneidae were dominants in numbers, while plexwassupplementedwith Oxyopesheterophthalmus^ Thomisidae comprised the main partofthe biomass Cheiracanthiumpennyi^Thomisusonustus^andRuncinia (Efimik 1989). According to our data, Araneidae grammica, and in the otherone (Solenoozerny) with was the most abundant familywhileTetragnathidae Pellenes seriatus and Yllenus vittatus. The presence occasionally occurred in steppe as rare finds. In the ofthe latter three species and the abundance ofH. east ofthe Russian Plain, in stony and forb steppes lineiventris is characteristic of the southern sandy ofSamarskaya Luka,Thomisidae reached the high- steppes. est abundance followed byAraneidae and Salticidae (KRASNOBAEV 2003). In eastern Hungary (Great Discussion Hungarian Plain), drysandygrassland communities Spiderspeciesrichnessoftheinvestigatedsteppefrag- that are close to sandy steppes were also dominated mentsdependedonthesteppetype.Thelownumber by Araneidae and Thomisidae while Salticidae and ofspecies on the largest site ofbunchgrass steppes Dictynidaecomprisedasecond-rankedgroup(HOR- canbe explainedbythe monotonous plainreliefand VATH et al. 2009). absenceofneighbouringintrazonalhabitats.Presum- NosinglespeciesdominatedinaUthesteppetypes ably, invertebrate communities within small sites and evenwithin one type,a dominance complexwas increase in species richness due to species exchange composedofdifferentspecies.Meadowsteppeswere with adjacent habitats (MAGURA & KODOBOCZ dominatedbywetlandspeciesandgenuinesteppesby 2007). This statement is true for the forest-steppe wide-spreadgrassland species.In azonalsteppes,the and northernpartofthe steppezonewhere an active complexincludedspeciesinhabitingsparsevegetation interchange takes place (CHERNOV 1975). In our ofsandyandchalkysoils.Dominantspiderspeciesof investigations,werecordedthepresenceofforestand steppe and steppe-like habitats oftheUrals alsovary wetland species in meadow steppes. However in the depending on zonal and local conditions (ESYUNIN south ofthe steppe zone, despite a rich local fauna 2009). in general and the existence ofneighbouring forest Inconclusionthespiderassemblagesofthenorth- stands and lakes, the species composition ofspiders ernmostmeadowsteppesandthesouthernmostsandy ofsteppe fragments remained the poorest. Onlyone steppes differeddramatically, as expected,because of wetland specieswas found there. zonal-climatic differences. From a zonal standpoint, About 12%ofthe species oftheinvestigatedarea thefirsthavemuchincommonwithmeadows(PON- & occurred in all steppe types. Another 12% changed OMAREV POLCHANINOVA 2006), and the latter their habitat preference spreading from north to withbrackish meadows and salt-marsheswith dense south and moved from steppe sites to shaded and vegetation(POLCHANINOVA1998).Inourstudy,the moist ones. These data illustrate well the ‘principle most specificwere herb dwelling spider assemblages ofzonal change ofhabitats’by BEI-BIENKO (1966). ofthebunchgrass stepperepresentedinUkrainebya Someofthem,suchasNeottiurabimaculata^changed single site in the middle ofplowed Dnieperlowland. 74 N.Polchaninovo Thesecondspecificgroupswereassemblagesofsandy GribovaS.A,T.I.Isachenko&Y.M.Lavrenko (eds) steppesinthenorthernsteppesubzonebecausethese (1980):[VegetationoftheEuropeanpartoftheUSSR]. fragmentsareisolatedanddifferinmicroclimaticcon- Nauka,Leningrad.432pp. [in Russian]. ditionsandvegetationstructurefromothergrasslands. Horvath R., T. Magura, C. Szinetar & B. Toth- Steppes of a certain type are formed under both MERESZ (2009): Spiders are not less diverse in small climate and soil conditions of the zone and local garnadssilsaonldast:edAgrfaiseslldansdtsu,dybu(tEalesstsHduivnegrasreyi,nNoyvierrsgerga).ze-d topography (GrIBOVA et al. 1980). Accordingly, the Agriculture, Ecosystems and Environment 130: 16-22 structure ofthe animal communities ofthe steppes -doi: 10.1016/j.agee.2008.11.011 is a reflection ofthese conditions (MORDKOVITCH HASERS.E.&N.J.HUNTLY(2006):Thebiogeographyof 1982). Assemblages of herb-dwelling spiders are smallmammalsoffragmentedsagebrush-steppelands- characteristicofeachsteppetypeanddifferinspecies capes. -Journal ofMammology 87: 1165-1174- doi: composition and dominance structure. 10.1644/05-MAMM-A-385R2.1 KotenkoT.I. (1996): [Steppes ofUkraine: their impor- Acknowledgements tance,modernstate,scientificvalue,protectionpriority]. The authorisgratefultoJ. Gergerichforrevisingthe first -Vestnikzooologii 1996(1/2): 10-26 draft ofthe English manuscript and to A. Atemasov for KRASNOBAEVYR(2003): [Structureofherpetobiontand the map design. hortobiont spider populations in Samarskaya Luka]. In: Izvestia Samarskogo nauchnogo tsentra Rossiyskoi References akademii nauk. Spetsialnyi vypusk 1. 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